PeerJ:Evolutionary Studieshttps://peerj.com/articles/index.atom?journal=peerj&subject=1500Evolutionary Studies articles published in PeerJTransitional evolutionary forms in chasmosaurine ceratopsid dinosaurs: evidence from the Campanian of New Mexicohttps://peerj.com/articles/92512020-06-052020-06-05Denver W. FowlerElizabeth A. Freedman Fowler
Three new chasmosaurines from the Kirtland Formation (~75.0–73.4 Ma), New Mexico, form morphological and stratigraphic intermediates between Pentaceratops (~74.7–75 Ma, Fruitland Formation, New Mexico) and Anchiceratops (~72–71 Ma, Horseshoe Canyon Formation, Alberta). The new specimens exhibit gradual enclosure of the parietal embayment that characterizes Pentaceratops, providing support for the phylogenetic hypothesis that Pentaceratops and Anchiceratops are closely related. This stepwise change of morphologic characters observed in chasmosaurine taxa that do not overlap stratigraphically is supportive of evolution by anagenesis. Recently published hypotheses that place Pentaceratops and Anchiceratops into separate clades are not supported. This phylogenetic relationship demonstrates unrestricted movement of large-bodied taxa between hitherto purported northern and southern provinces in the late Campanian, weakening support for the hypothesis of extreme faunal provincialism in the Late Cretaceous Western Interior.

Three new chasmosaurines from the Kirtland Formation (~75.0–73.4 Ma), New Mexico, form morphological and stratigraphic intermediates between Pentaceratops (~74.7–75 Ma, Fruitland Formation, New Mexico) and Anchiceratops (~72–71 Ma, Horseshoe Canyon Formation, Alberta). The new specimens exhibit gradual enclosure of the parietal embayment that characterizes Pentaceratops, providing support for the phylogenetic hypothesis that Pentaceratops and Anchiceratops are closely related. This stepwise change of morphologic characters observed in chasmosaurine taxa that do not overlap stratigraphically is supportive of evolution by anagenesis. Recently published hypotheses that place Pentaceratops and Anchiceratops into separate clades are not supported. This phylogenetic relationship demonstrates unrestricted movement of large-bodied taxa between hitherto purported northern and southern provinces in the late Campanian, weakening support for the hypothesis of extreme faunal provincialism in the Late Cretaceous Western Interior.

A high-resolution growth series of Tyrannosaurus rex obtained from multiple lines of evidencehttps://peerj.com/articles/91922020-06-042020-06-04Thomas D. Carr
Background
During the growth of complex multicellular organisms, chronological age, size and morphology change together in a hierarchical and coordinated pattern. Among extinct species, the growth of Tyrannosaurus rex has received repeated attention through quantitative analyses of relative maturity and chronological age. Its growth series shows an extreme transformation from shallow skulls in juveniles to deep skulls in adults along with a reduction in tooth count, and its growth curve shows that T. rex had a high growth rate in contrast to its closest relatives. However, separately, these sets of data provide an incomplete picture of the congruence between age, size, and relative maturity in this exemplar species. The goal of this work is to analyze these data sets together using cladistic analysis to produce a single hypothesis of growth that includes all of the relevant data.
Methods
The three axes of growth were analyzed together using cladistic analysis, based on a data set of 1,850 morphological characters and 44 specimens. The analysis was run in TNT v.1.5 under a New Technology search followed by a Traditional search. Correlation tests were run in IBM SPSS Statistics v. 24.0.0.0.
Results
An initial analysis that included all of the specimens recovered 50 multiple most parsimonious ontograms a series of analyses identified 13 wildcard specimens. An analysis run without the wildcard specimens recovered a single most parsimonious tree (i.e., ontogram) of 3,053 steps. The ontogram is composed of 21 growth stages, and all but the first and third are supported by unambiguously optimized synontomorphies. T. rex ontogeny can be divided into five discrete growth categories that are diagnosed by chronological age, morphology, and, in part, size (uninformative among adults). The topology shows that the transition from shallow to deep skull shape occurred between 13 and 15 years of age, and the size of the immediate relatives of T. rex was exceeded between its 15th and 18th years. Although size and maturity are congruent among juveniles and subadults, congruence is not seen among adults; for example, one of the least mature adults (RSM 2523.8) is also the largest and most massive example of the species. The extreme number of changes at the transition between juveniles and subadults shows that the ontogeny of T. rex exhibits secondary metamorphosis, analogous to the abrupt ontogenetic changes that are seen at sexual maturity among teleosts. These results provide a point of comparison for testing the congruence between maturity and chronological age, size, and mass, as well as integrating previous work on functional morphology into a rigorous ontogenetic framework. Comparison of the growth series of T. rex with those of outgroup taxa clarifies the ontogenetic trends that were inherited from the common ancestor of Archosauriformes.

Background

During the growth of complex multicellular organisms, chronological age, size and morphology change together in a hierarchical and coordinated pattern. Among extinct species, the growth of Tyrannosaurus rex has received repeated attention through quantitative analyses of relative maturity and chronological age. Its growth series shows an extreme transformation from shallow skulls in juveniles to deep skulls in adults along with a reduction in tooth count, and its growth curve shows that T. rex had a high growth rate in contrast to its closest relatives. However, separately, these sets of data provide an incomplete picture of the congruence between age, size, and relative maturity in this exemplar species. The goal of this work is to analyze these data sets together using cladistic analysis to produce a single hypothesis of growth that includes all of the relevant data.

Methods

The three axes of growth were analyzed together using cladistic analysis, based on a data set of 1,850 morphological characters and 44 specimens. The analysis was run in TNT v.1.5 under a New Technology search followed by a Traditional search. Correlation tests were run in IBM SPSS Statistics v. 24.0.0.0.

Results

An initial analysis that included all of the specimens recovered 50 multiple most parsimonious ontograms a series of analyses identified 13 wildcard specimens. An analysis run without the wildcard specimens recovered a single most parsimonious tree (i.e., ontogram) of 3,053 steps. The ontogram is composed of 21 growth stages, and all but the first and third are supported by unambiguously optimized synontomorphies. T. rex ontogeny can be divided into five discrete growth categories that are diagnosed by chronological age, morphology, and, in part, size (uninformative among adults). The topology shows that the transition from shallow to deep skull shape occurred between 13 and 15 years of age, and the size of the immediate relatives of T. rex was exceeded between its 15th and 18th years. Although size and maturity are congruent among juveniles and subadults, congruence is not seen among adults; for example, one of the least mature adults (RSM 2523.8) is also the largest and most massive example of the species. The extreme number of changes at the transition between juveniles and subadults shows that the ontogeny of T. rex exhibits secondary metamorphosis, analogous to the abrupt ontogenetic changes that are seen at sexual maturity among teleosts. These results provide a point of comparison for testing the congruence between maturity and chronological age, size, and mass, as well as integrating previous work on functional morphology into a rigorous ontogenetic framework. Comparison of the growth series of T. rex with those of outgroup taxa clarifies the ontogenetic trends that were inherited from the common ancestor of Archosauriformes.

New insights into the giant mustelids (Mammalia, Carnivora, Mustelidae) from Langebaanweg fossil site (West Coast Fossil Park, South Africa, early Pliocene)https://peerj.com/articles/92212020-06-012020-06-01Alberto ValencianoRomala Govender
Giant mustelids are a paraphyletic group of mustelids found in the Neogene of Eurasia, Africa and North America. Most are known largely from dental remains, with their postcranial skeleton mostly unknown. Here, we describe new craniodental and postcranial remains of the large lutrine Sivaonyx hendeyi and the leopard-size gulonine Plesiogulo aff. monspessulanus from the early Pliocene site Langebaanweg, South Africa. The new material of the endemic S. hendeyi, includes upper incisors and premolars, and fragmentary humerus, ulna and a complete astragalus. Its postcrania shares more traits with the living Aonyx capensis than the late Miocene Sivaonyx beyi from Chad. Sivaonyx hendeyi could therefore be tentatively interpreted as a relatively more aquatic taxon than the Chadian species, comparable to A. capensis. The new specimens of Plesiogulo comprise two edentulous maxillae, including one of a juvenile individual with incomplete decidual dentition, and a fragmentary forelimb of an adult individual. The new dental measurements point to this form being amongst the largest specimens of the genus. Both P3-4 differs from the very large species Plesiogulo botori from late Miocene of Kenya and Ethiopia. This confirms the existence of two distinct large species of Plesiogulo in Africa during the Mio/Pliocene, P. botori in the Late Miocene of Eastern Africa (6.1–5.5 Ma) and Plesiogulo aff. monspessulanus at the beginning of the Pliocene in southern Africa (5.2 Ma). Lastly, we report for the first time the presence of both Sivaonyx and Plesiogulo in MPPM and LQSM at Langebaanweg, suggesting that the differences observed from the locality may be produced by sedimentation or sampling biases instead of temporal replacement within the carnivoran guild.

Giant mustelids are a paraphyletic group of mustelids found in the Neogene of Eurasia, Africa and North America. Most are known largely from dental remains, with their postcranial skeleton mostly unknown. Here, we describe new craniodental and postcranial remains of the large lutrine Sivaonyx hendeyi and the leopard-size gulonine Plesiogulo aff. monspessulanus from the early Pliocene site Langebaanweg, South Africa. The new material of the endemic S. hendeyi, includes upper incisors and premolars, and fragmentary humerus, ulna and a complete astragalus. Its postcrania shares more traits with the living Aonyx capensis than the late Miocene Sivaonyx beyi from Chad. Sivaonyx hendeyi could therefore be tentatively interpreted as a relatively more aquatic taxon than the Chadian species, comparable to A. capensis. The new specimens of Plesiogulo comprise two edentulous maxillae, including one of a juvenile individual with incomplete decidual dentition, and a fragmentary forelimb of an adult individual. The new dental measurements point to this form being amongst the largest specimens of the genus. Both P3-4 differs from the very large species Plesiogulo botori from late Miocene of Kenya and Ethiopia. This confirms the existence of two distinct large species of Plesiogulo in Africa during the Mio/Pliocene, P. botori in the Late Miocene of Eastern Africa (6.1–5.5 Ma) and Plesiogulo aff. monspessulanus at the beginning of the Pliocene in southern Africa (5.2 Ma). Lastly, we report for the first time the presence of both Sivaonyx and Plesiogulo in MPPM and LQSM at Langebaanweg, suggesting that the differences observed from the locality may be produced by sedimentation or sampling biases instead of temporal replacement within the carnivoran guild.

Characterization of the bacterial microbiota composition and evolution at different intestinal tract in wild pigs (Sus scrofa ussuricus)https://peerj.com/articles/91242020-05-262020-05-26Guangli YangChuanxin ShiShuhong ZhangYan LiuZhiqiang LiFengyi GaoYanyan CuiYongfeng YanMing Li
Commensal microorganisms are essential to the normal development and function of many aspects of animal biology, including digestion, nutrient absorption, immunological development, behaviors, and evolution. The specific microbial composition and evolution of the intestinal tracts of wild pigs remain poorly characterized. This study therefore sought to assess the composition, distribution, and evolution of the intestinal microbiome of wild pigs. For these analyses, 16S rRNA V3-V4 regions from five gut sections prepared from each of three wild sows were sequenced to detect the microbiome composition. These analyses revealed the presence of 6,513 operational taxonomic units (OTUs) mostly distributed across 17 phyla and 163 genera in these samples, with Firmicutes and Actinobacteria being the most prevalent phyla of microbes present in cecum and jejunum samples, respectively. Moreover, the abundance of Actinobacteria in wild pigs was higher than that in domestic pigs. At the genus level the Bifidobacterium and Allobaculum species of microbes were most abundant in all tested gut sections, with higher relative abundance in wild pigs relative to domestic pigs, indicating that in the process of pig evolution, the intestinal microbes also evolved, and changes in the intestinal microbial diversity could have been one of the evolutionary forces of pigs. Intestinal microbial functional analyses also revealed the microbes present in the small intestine (duodenum, jejunum, and ileum) and large intestine (cecum and colon) of wild pigs to engage distinct metabolic spatial structures and pathways relative to one another. Overall, these results offer unique insights that would help to advance the current understanding of how the intestinal microbes interact with the host and affect the evolution of pigs.

Commensal microorganisms are essential to the normal development and function of many aspects of animal biology, including digestion, nutrient absorption, immunological development, behaviors, and evolution. The specific microbial composition and evolution of the intestinal tracts of wild pigs remain poorly characterized. This study therefore sought to assess the composition, distribution, and evolution of the intestinal microbiome of wild pigs. For these analyses, 16S rRNA V3-V4 regions from five gut sections prepared from each of three wild sows were sequenced to detect the microbiome composition. These analyses revealed the presence of 6,513 operational taxonomic units (OTUs) mostly distributed across 17 phyla and 163 genera in these samples, with Firmicutes and Actinobacteria being the most prevalent phyla of microbes present in cecum and jejunum samples, respectively. Moreover, the abundance of Actinobacteria in wild pigs was higher than that in domestic pigs. At the genus level the Bifidobacterium and Allobaculum species of microbes were most abundant in all tested gut sections, with higher relative abundance in wild pigs relative to domestic pigs, indicating that in the process of pig evolution, the intestinal microbes also evolved, and changes in the intestinal microbial diversity could have been one of the evolutionary forces of pigs. Intestinal microbial functional analyses also revealed the microbes present in the small intestine (duodenum, jejunum, and ileum) and large intestine (cecum and colon) of wild pigs to engage distinct metabolic spatial structures and pathways relative to one another. Overall, these results offer unique insights that would help to advance the current understanding of how the intestinal microbes interact with the host and affect the evolution of pigs.

Comparative analysis of four Zantedeschia chloroplast genomes: expansion and contraction of the IR region, phylogenetic analyses and SSR genetic diversity assessmenthttps://peerj.com/articles/91322020-05-222020-05-22Shuilian HeYang YangZiwei LiXuejiao WangYanbing GuoHongzhi Wu
The horticulturally important genus Zantedeschia (Araceae) comprises eight species of herbaceous perennials. We sequenced, assembled and analyzed the chloroplast (cp) genomes of four species of Zantedeschia (Z. aethiopica, Z. odorata, Z. elliottiana, and Z. rehmannii) to investigate the structure of the cp genome in the genus. According to our results, the cp genome of Zantedeschia ranges in size from 169,065 bp (Z. aethiopica) to 175,906 bp (Z. elliottiana). We identified a total of 112 unique genes, including 78 protein-coding genes, 30 transfer RNA (tRNA) genes and four ribosomal RNA (rRNA) genes. Comparison of our results with cp genomes from other species in the Araceae suggests that the relatively large sizes of the Zantedeschia cp genomes may result from inverted repeats (IR) region expansion. The sampled Zantedeschia species formed a monophylogenetic clade in our phylogenetic analysis. Furthermore, the long single copy (LSC) and short single copy (SSC) regions in Zantedeschia are more divergent than the IR regions in the same genus, and non-coding regions showed generally higher divergence than coding regions. We identified a total of 410 cpSSR sites from the four Zantedeschia species studied. Genetic diversity analyses based on four polymorphic SSR markers from 134 cultivars of Zantedeschia suggested that high genetic diversity (I = 0.934; Ne = 2.371) is present in the Zantedeschia cultivars. High genetic polymorphism from the cpSSR region suggests that cpSSR could be an effective tool for genetic diversity assessment and identification of Zantedeschia varieties.

The horticulturally important genus Zantedeschia (Araceae) comprises eight species of herbaceous perennials. We sequenced, assembled and analyzed the chloroplast (cp) genomes of four species of Zantedeschia (Z. aethiopica, Z. odorata, Z. elliottiana, and Z. rehmannii) to investigate the structure of the cp genome in the genus. According to our results, the cp genome of Zantedeschia ranges in size from 169,065 bp (Z. aethiopica) to 175,906 bp (Z. elliottiana). We identified a total of 112 unique genes, including 78 protein-coding genes, 30 transfer RNA (tRNA) genes and four ribosomal RNA (rRNA) genes. Comparison of our results with cp genomes from other species in the Araceae suggests that the relatively large sizes of the Zantedeschia cp genomes may result from inverted repeats (IR) region expansion. The sampled Zantedeschia species formed a monophylogenetic clade in our phylogenetic analysis. Furthermore, the long single copy (LSC) and short single copy (SSC) regions in Zantedeschia are more divergent than the IR regions in the same genus, and non-coding regions showed generally higher divergence than coding regions. We identified a total of 410 cpSSR sites from the four Zantedeschia species studied. Genetic diversity analyses based on four polymorphic SSR markers from 134 cultivars of Zantedeschia suggested that high genetic diversity (I = 0.934; Ne = 2.371) is present in the Zantedeschia cultivars. High genetic polymorphism from the cpSSR region suggests that cpSSR could be an effective tool for genetic diversity assessment and identification of Zantedeschia varieties.

Endosymbiotic adaptations in three new bacterial species associated with Dictyostelium discoideum: Paraburkholderia agricolaris sp. nov., Paraburkholderia hayleyella sp. nov., and Paraburkholderia bonniea sp. novhttps://peerj.com/articles/91512020-05-222020-05-22Debra A. BrockSuegene NohAlicia N.M. HubertTamara S. HaselkornSusanne DiSalvoMelanie K. SuessAlexander S. BradleyMahboubeh Tavakoli-NezhadKatherine S. GeistDavid C. QuellerJoan E. Strassmann
Here we give names to three new species of Paraburkholderia that can remain in symbiosis indefinitely in the spores of a soil dwelling eukaryote, Dictyostelium discoideum. The new species P. agricolaris sp. nov., P. hayleyella sp. nov., and P. bonniea sp. nov. are widespread across the eastern USA and were isolated as internal symbionts of wild-collected D. discoideum. We describe these sp. nov. using several approaches. Evidence that they are each a distinct new species comes from their phylogenetic position, average nucleotide identity, genome-genome distance, carbon usage, reduced length, cooler optimal growth temperature, metabolic tests, and their previously described ability to invade D. discoideum amoebae and form a symbiotic relationship. All three of these new species facilitate the prolonged carriage of food bacteria by D. discoideum, though they themselves are not food. Further studies of the interactions of these three new species with D. discoideum should be fruitful for understanding the ecology and evolution of symbioses.

Here we give names to three new species of Paraburkholderia that can remain in symbiosis indefinitely in the spores of a soil dwelling eukaryote, Dictyostelium discoideum. The new species P. agricolaris sp. nov., P. hayleyella sp. nov., and P. bonniea sp. nov. are widespread across the eastern USA and were isolated as internal symbionts of wild-collected D. discoideum. We describe these sp. nov. using several approaches. Evidence that they are each a distinct new species comes from their phylogenetic position, average nucleotide identity, genome-genome distance, carbon usage, reduced length, cooler optimal growth temperature, metabolic tests, and their previously described ability to invade D. discoideum amoebae and form a symbiotic relationship. All three of these new species facilitate the prolonged carriage of food bacteria by D. discoideum, though they themselves are not food. Further studies of the interactions of these three new species with D. discoideum should be fruitful for understanding the ecology and evolution of symbioses.

The effect of landscape on functional connectivity and shell shape in the land snail Humboldtiana durangoensishttps://peerj.com/articles/91772020-05-202020-05-20Benjamín LópezOmar MejíaGerardo Zúñiga
The populations of Humboldtiana durangoensis have experienced a drastic reduction in the effective population size; in addition, the species is threatened by anthropogenic activities. For the aforementioned, landscape genetics will serve as a tool to define the potential evolutionarily significant units (ESU) for this species. To complete our objective, we evaluated the effect of cover vegetation and climate on the functional connectivity of the species from the last glacial maximum (LGM) to the present as well as the effect of climate on shell shape. Partial Mantel tests, distance-based redundance analysis and a Bayesian framework were used to evaluate connectivity. On the other hand, geometric morphometrics, phylogenetic principal component analysis and redundancy analysis were used for the analysis of shell shape. Our results suggest that the suitable areas have been decreasing since the LGM; also, vegetation cover rather than climate has influenced the genetic connectivity among land snail populations, although temperature had a high influence on shell shape in this species. In conclusion, vegetation cover was the main factor that determined the functional connectivity for the land snail; however, local selective pressures led to different phenotypes in shell shape that allowed us to postulate that each one of the previously defined genetic groups must be considered as a different ESU.

The populations of Humboldtiana durangoensis have experienced a drastic reduction in the effective population size; in addition, the species is threatened by anthropogenic activities. For the aforementioned, landscape genetics will serve as a tool to define the potential evolutionarily significant units (ESU) for this species. To complete our objective, we evaluated the effect of cover vegetation and climate on the functional connectivity of the species from the last glacial maximum (LGM) to the present as well as the effect of climate on shell shape. Partial Mantel tests, distance-based redundance analysis and a Bayesian framework were used to evaluate connectivity. On the other hand, geometric morphometrics, phylogenetic principal component analysis and redundancy analysis were used for the analysis of shell shape. Our results suggest that the suitable areas have been decreasing since the LGM; also, vegetation cover rather than climate has influenced the genetic connectivity among land snail populations, although temperature had a high influence on shell shape in this species. In conclusion, vegetation cover was the main factor that determined the functional connectivity for the land snail; however, local selective pressures led to different phenotypes in shell shape that allowed us to postulate that each one of the previously defined genetic groups must be considered as a different ESU.

The case for extended lifespan in cooperatively breeding mammals: a re-appraisalhttps://peerj.com/articles/92142020-05-192020-05-19Jack Thorley
Recent comparative studies have suggested that cooperative breeding is associated with increases in maximum lifespan among mammals, replicating a pattern also seen in birds and insects. In this study, we re-examine the case for increased lifespan in mammalian cooperative breeders by analysing a large dataset of maximum longevity records. We did not find any consistent, strong evidence that cooperative breeders have longer lifespans than other mammals after having controlled for variation in body mass, mode of life and data quality. The only possible exception to this general trend is found in the African mole-rats (the Bathyergid family), where all members are relatively long-lived, but where the social, cooperatively breeding species appear to be much longer-lived than the solitary species. However, solitary mole-rat species have rarely been kept in captivity or followed longitudinally in the wild and so it seems likely that their maximum lifespan has been underestimated when compared to the highly researched social species. Although few subterranean mammals have received much attention in a captive or wild setting, current data instead supports a causal role of subterranean living on lifespan extension in mammals.

Recent comparative studies have suggested that cooperative breeding is associated with increases in maximum lifespan among mammals, replicating a pattern also seen in birds and insects. In this study, we re-examine the case for increased lifespan in mammalian cooperative breeders by analysing a large dataset of maximum longevity records. We did not find any consistent, strong evidence that cooperative breeders have longer lifespans than other mammals after having controlled for variation in body mass, mode of life and data quality. The only possible exception to this general trend is found in the African mole-rats (the Bathyergid family), where all members are relatively long-lived, but where the social, cooperatively breeding species appear to be much longer-lived than the solitary species. However, solitary mole-rat species have rarely been kept in captivity or followed longitudinally in the wild and so it seems likely that their maximum lifespan has been underestimated when compared to the highly researched social species. Although few subterranean mammals have received much attention in a captive or wild setting, current data instead supports a causal role of subterranean living on lifespan extension in mammals.

The mitochondrial genome of Acrobeloides varius (Cephalobomorpha) confirms non-monophyly of Tylenchina (Nematoda)https://peerj.com/articles/91082020-05-132020-05-13Taeho KimYucheol LeeHyun-Jong KilJoong-Ki Park
The infraorder Cephalobomorpha is a diverse and ecologically important nematode group found in almost all terrestrial environments. In a recent nematode classification system based on SSU rDNA, Cephalobomorpha was classified within the suborder Tylenchina with Panagrolaimomorpha, Tylenchomorpha and Drilonematomorpha. However, phylogenetic relationships among species within Tylenchina are not always consistent, and the phylogenetic position of Cephalobomorpha is still uncertain. In this study, in order to examine phylogenetic relationships of Cephalobomorpha with other nematode groups, we determined the complete mitochondrial genome sequence of Acrobeloides varius, the first sequenced representative of Cephalobomorpha, and used this sequence for phylogenetic analyses along with 101 other nematode species. Phylogenetic analyses using amino acid and nucleotide sequence data of 12 protein-coding genes strongly support a sister relationship between the two cephalobomorpha species A. varius and Acrobeles complexus (represented by a partial mt genome sequence). In this mitochondrial genome phylogeny, Cephalobomorpha was sister to all chromadorean species (excluding Plectus acuminatus of Plectida) and separated from Panagrolaimomorpha and Tylenchomorpha, rendering Tylenchina non-monophyletic. Mitochondrial gene order among Tylenchina species is not conserved, and gene clusters shared between A. varius and A. complexus are very limited. Results from phylogenetic analysis and gene order comparison confirms Tylenchina is not monophyletic. To better understand phylogenetic relationships among Tylenchina members, additional mitochondrial genome information is needed from underrepresented taxa representing Panagrolaimomorpha and Cephalobomorpha.

The infraorder Cephalobomorpha is a diverse and ecologically important nematode group found in almost all terrestrial environments. In a recent nematode classification system based on SSU rDNA, Cephalobomorpha was classified within the suborder Tylenchina with Panagrolaimomorpha, Tylenchomorpha and Drilonematomorpha. However, phylogenetic relationships among species within Tylenchina are not always consistent, and the phylogenetic position of Cephalobomorpha is still uncertain. In this study, in order to examine phylogenetic relationships of Cephalobomorpha with other nematode groups, we determined the complete mitochondrial genome sequence of Acrobeloides varius, the first sequenced representative of Cephalobomorpha, and used this sequence for phylogenetic analyses along with 101 other nematode species. Phylogenetic analyses using amino acid and nucleotide sequence data of 12 protein-coding genes strongly support a sister relationship between the two cephalobomorpha species A. varius and Acrobeles complexus (represented by a partial mt genome sequence). In this mitochondrial genome phylogeny, Cephalobomorpha was sister to all chromadorean species (excluding Plectus acuminatus of Plectida) and separated from Panagrolaimomorpha and Tylenchomorpha, rendering Tylenchina non-monophyletic. Mitochondrial gene order among Tylenchina species is not conserved, and gene clusters shared between A. varius and A. complexus are very limited. Results from phylogenetic analysis and gene order comparison confirms Tylenchina is not monophyletic. To better understand phylogenetic relationships among Tylenchina members, additional mitochondrial genome information is needed from underrepresented taxa representing Panagrolaimomorpha and Cephalobomorpha.

Thecodont tooth attachment and replacement in bolosaurid parareptileshttps://peerj.com/articles/91682020-05-132020-05-13Adam J. SnyderAaron R.H. LeBlancChen JunJoseph J. BevittRobert R. Reisz
Permian bolosaurid parareptiles are well-known for having complex tooth crowns and complete tooth rows in the jaws, in contrast to the comparatively simple teeth and frequent replacement gaps in all other Paleozoic amniotes. Analysis of the specialized dentition of the bolosaurid parareptiles Bolosaurus from North America and Belebey from Russia, utilizing a combination of histological and tomographic data, reveals unusual patterns of tooth development and replacement. The data confirm that bolosaurid teeth have thecodont implantation with deep roots, the oldest known such example among amniotes, and independently evolved among much younger archosauromorphs (including dinosaurs and crocodilians) and among synapsids (including mammals). High-resolution CT scans were able to detect the density boundary between the alveolar bone and the jawbone, as confirmed by histology, and revealed the location and size of developing replacement teeth in the pulp cavity of functional teeth. Evidence provided by the paratype dentary of Belebey chengi indicates that replacement teeth are present along the whole tooth row at slightly different stages of development, with the ontogenetically more developed teeth anteriorly, suggesting that tooth replacement was highly synchronized. CT data also show tooth replacement is directly related to the presence of lingual pits in the jaw, and that migration of tooth buds occurs initially close to these resorption pits to a position immediately below the functional tooth within its pulp cavity. The size and complex shape of the replacement teeth in the holotype of Bolosaurus grandis indicate that the replacement teeth can develop within the pulp cavity to an advanced stage while the previous generation remains functional for an extended time, reminiscent of the condition seen in other amniotes with occluding dentitions, including mammals.

Permian bolosaurid parareptiles are well-known for having complex tooth crowns and complete tooth rows in the jaws, in contrast to the comparatively simple teeth and frequent replacement gaps in all other Paleozoic amniotes. Analysis of the specialized dentition of the bolosaurid parareptiles Bolosaurus from North America and Belebey from Russia, utilizing a combination of histological and tomographic data, reveals unusual patterns of tooth development and replacement. The data confirm that bolosaurid teeth have thecodont implantation with deep roots, the oldest known such example among amniotes, and independently evolved among much younger archosauromorphs (including dinosaurs and crocodilians) and among synapsids (including mammals). High-resolution CT scans were able to detect the density boundary between the alveolar bone and the jawbone, as confirmed by histology, and revealed the location and size of developing replacement teeth in the pulp cavity of functional teeth. Evidence provided by the paratype dentary of Belebey chengi indicates that replacement teeth are present along the whole tooth row at slightly different stages of development, with the ontogenetically more developed teeth anteriorly, suggesting that tooth replacement was highly synchronized. CT data also show tooth replacement is directly related to the presence of lingual pits in the jaw, and that migration of tooth buds occurs initially close to these resorption pits to a position immediately below the functional tooth within its pulp cavity. The size and complex shape of the replacement teeth in the holotype of Bolosaurus grandis indicate that the replacement teeth can develop within the pulp cavity to an advanced stage while the previous generation remains functional for an extended time, reminiscent of the condition seen in other amniotes with occluding dentitions, including mammals.